Graduate Theses & Dissertations

cascading effects of risk in the wild
Predation risk can elicit a range of responses in prey, but to date little is known about breadth of potential responses that may arise under realistic field conditions and how such responses are linked, leaving a fragmented picture of risk-related consequences on individuals. We increased predation risk in free-ranging snowshoe hares (Lepus americanus) during two consecutive summers by simulating natural chases using a model predator (i.e., domestic dog), and monitored hare stress physiology, energy expenditure, behaviour, condition, and habitat use. We show that higher levels of risk elicited marked changes in physiological stress metrics including sustained high levels of free plasma cortisol which had cascading effects on glucose, and immunology, but not condition. Risk-augmented hares also had lowered daily energy expenditure, spent more time foraging, and decreased rest, vigilance, and travel. It is possible that these alterations allowed risk-exposed hares to increase their condition at the same rate as controls. Additionally, risk-augmented hares selected, had high fidelity to, and were more mobile in structurally dense habitat (i.e., shrubs) which provided them additional cover from predators. They also used more open habitat (i.e., conifer) differently based on locale within the home range, using familiar conifer areas within cores for rest while moving through unfamiliar conifer areas in the periphery. Overall, these findings show that prey can have a multi-faceted, highly plastic response in the face of risk and can mitigate the effects of their stress physiology given the right environmental conditions. Author Keywords: behaviour, condition, daily energy expenditure, predator-prey interactions, snowshoe hare, stress physiology
Habitat Characteristics, Density Patterns and Environmental Niches of Indo-Pacific Humpback Dolphins (Sousa chinensis) of the Pearl River Estuary and Eastern Taiwan Strait
The purpose of this thesis is to quantify the habitat characteristics, density patterns and environmental niches of two groups of Indo-Pacific humpback dolphins: Chinese white dolphins (CWD) of the Pearl River estuary (PRE), and Taiwanese white dolphins (TWD, =Taiwanese humpback dolphin, THD) found in the eastern Taiwan Strait (ETS). Much work has already been done on the habitat use of CWDs in parts of the PRE, so the purpose of my first two chapters was to advance knowledge of the TWD to a comparable level. Chapter 2 contains the first published description of the relatively shallow, inshore, estuarine habitat of the TWD. General environmental characteristics and observed group sizes were consistent with other populations of humpback dolphins, and group sizes were not correlated with the environmental variables measured during surveys. Chapter 3 investigated density patterns of TWDs, finding spatiotemporal heterogeneity across the study area. Humpback dolphin densities fluctuated from year to year, but some parts of the study area were consistently used more than others. Environmental characteristics again did not influence dolphin densities, though more dolphins than expected were sighted in waters adjacent to major land reclamations, which may be related to the location of these areas close to major rivers. In Chapter 4, niches of the TWD and CWDs found in the PRE were compared using species distribution models, which indicated significant niche overlap. This may be due to niche conservatism maintaining similar fundamental niches between the two groups since their historical split >10,000 years ago, or a result of the intrinsic biotic factors that influence occurrence data affecting the hypervolume dimensions of each realized niche in similar ways. Geographic predictions indicate that most of the TWD’s range has likely been surveyed, and that there may be connectivity between PRE humpback dolphins and at least one neighbouring putative population due to continuous predicted suitable habitat in waters that remain poorly surveyed. Overall, my thesis demonstrates that density patterns may vary over time, but on a broad temporal scale, these two allopatric groups of Indo-Pacific humpback dolphins have similar habitat requirements in geographically isolated, but environmentally similar locations. Author Keywords: density, habitat, Indo-Pacific humpback dolphin, niche overlap, Sousa chinensis, species distribution model
New Interpretations from Old Data
Range contractions and expansions are important ecological concepts for species management decisions. These decisions relate not only to rare and endangered species but to common and invasive species as well. The development of the broad spatiotemporal extent models that are helpful in examining range fluctuations can be challenging given the lack of data expansive enough to cover the time periods and geographic extents needed to fit the models. Archival records such as museum databases and harvest data can provide the spatiotemporal extent needed but present statistical challenges given they represent presence-only location information. In this thesis, I used maximum entropy and Bayesian hierarchical occupancy algorithms fitted with archival presence-only records to develop spatiotemporal models covering broad spatial and temporal extents for snowshoe hare and Canada lynx. These two algorithm types are well suited for presence-only data records and can be adapted to include biological and physical processes, thus improving the ecological realism of the models. Using these modelling methods, I found the extent of occurrence (EOO) and area of occupancy (AOO) varied greatly over time and space for both snowshoe hare and Canada lynx, suggesting that management decisions for these species should include consideration of these variations. While the presence-only data were appropriate for model development and understanding changing values in EOO and AOO, it sometimes lacked the locational accuracy and precision needed to create fine scale ecological analyses, thus resulting in somewhat coarse but potentially relevant conclusions. Author Keywords: Area of occupancy, Bayesian hierarchical models, Canada lynx, Extent of occurrence, Presence-only data, Snowshoe hare
Time to adapt
To better understand species’ resilience to climate change and implement solutions, we must conserve environments that maintain standing adaptive genetic variation and the potential generation of new beneficial alleles. Coding trinucleotide repeats (cTNRs) providing high-pace adaptive capabilities via high rates of mutation are ideal targets for mitigating the decline of species at risk by characterizing adaptively significant populations. Ultimately, adaptive genetic information will inform the protection of biological diversity below the species level (i.e., “Evolutionarily Significant Units” or “ESUs”). This dissertation investigates cTNRs within candidate genes to determine their prevalence and influence under selection in North American mammals. First, I evaluated the potential for somatic mosaicism in Canada lynx (Lynx canadensis), and found that tissue-specific mosaicism does not confound cTNR genotyping success in lynx. Second, I assessed a selection of clock gene cTNRs across characterized mammals and found that these repeats are abundant and highly variable in length and purity. I also identified preliminary signatures of selection in 3 clock gene cTNRs in 3 pairs of congeneric North American mammal species, highlighting the importance of cTNRs for understanding the evolution and adaptation of wild populations. I further evaluated the influence of selection on the NR1D1 cTNR within Canada lynx sampled across Canada using environmental correlation, where I estimated the variation in NR1D1 cTNR alleles explained by environmental and spatial variables after removing the effects of neutral population structure. Although most variation was explained by neutral structure, environment and spatial patterns in eastern lynx populations significantly explained some of the variation in NR1D1 alleles. To examine the role of island populations in the generation and distribution of adaptive genetic variation, I used 14 neutral microsatellites and a dinucleotide repeat within a gene linked to mammalian body size, IGF-1, and found that both genetic drift and natural selection influence the observed genetic diversity of insular lynx. Finally, I estimated the divergence dates of peripheral lynx populations and made recommendations towards the conservation of Canada lynx; high levels of genetic differentiation coupled with post-glacial colonization histories and patterns of divergence at cTNR loci suggest at least 4 ESUs for Canada lynx across their range. Author Keywords: adaptation, Canada lynx, candidate genes, coding trinucleotide repeats, evolution, natural selection
Beyond Habitat
My objective was to understand how individual variation, in conjunction with variation in habitat, can affect individual and population-level variation in animal space use. I used coyotes (Canis latrans) as a model species to investigate the roles of hybridization, an inherited intrinsic factor, and spatial memory, a learned intrinsic factor, on space use. I used a diversity of methods and approaches, including meta-regression, multiple imputation, simulations, resource selection functions, step selection functions, net-squared displacement analysis, and survival analysis. A major contribution was my investigation of the performance of multiple imputation in a meta-regression framework in Chapter 2. My simulations indicated that multiple imputation performs well in estimating missing data within a meta-regression framework in most situations. In Chapter 3, I used published studies of coyote home range size in a meta-regression analysis with multiple imputation to examine the relative roles of hybridization and environmental variables on coyote home range size across North America. I found that hybridization with Canis species was a leading factor driving variation in coyote space use at a continental scale. In Chapter 4, I used telemetry data for 62 coyotes in Newfoundland, Canada, to investigate the influence of cognitive maps on resource use. I found that resident coyotes used spatial memory of the landscape to select or avoid resources at spatial scales beyond their immediate sensory perception relative to transient coyotes, presumably increasing their fitness. Taken together, my dissertation demonstrates that intrinsic factors, such as genetic ancestry and spatial memory, can have substantial influences on how animals use space at both individual and population levels, and at both a local and a continental scales. Author Keywords: canis latrans, hybridization, meta-regression, multiple imputation, Newfoundland, spatial memory
Elemental Variation in Daphnia
Environmental variation can affect consumer trait expression and alter ecological and evolutionary dynamics in natural populations. However, although dietary nutrient content can vary by an order of magnitude in natural ecosystems, intra-specific differences in consumer responses to food quality have not been thoroughly investigated. Therefore, the purpose of my dissertation was to examine the influence of dietary nutrition and other environmental factors on consumer phenotypic variation using the freshwater cladoceran Daphnia. I conducted a series of complementary laboratory and field studies where I examined the effects of dietary phosphorus (P) content and additional biological/environmental variables (multi-elemental limitation, genetic variation, and temperature) on daphnid life-history, biochemistry, body elemental composition, and population growth. In general, phenotypic expression within a species varied significantly in response to all experimental variables, but the relative influence of each was highly context dependent. In my first chapter, I found that dietary P content and environmental calcium (Ca) concentrations both altered Daphnia body Ca:P ratios and growth rates of individuals and affected intrinsic rates of increase at the population level. However, food quality appeared to have a much larger effect on trait expression, and body Ca:P ratios were highly sensitive to other forms of dietary nutrient limitation. Next, I documented significant quantitative genetic variation and phenotypic plasticity in daphnid P content, growth, and P use efficiency of field collected animals grown across dietary P gradients. Trait expression was also influenced by genotype X diet interactions suggesting that consumer responses to dietary nutrient limitation can be heritable and may be adaptive in different nutrient environments. Finally, I found that temperature appeared to override food quality effects and decouple P metabolism in natural Daphnia populations, but total biomass production was affected by both dietary P content and temperature, depending on the nutrient content of the lake. Overall, my dissertation shows that consumer responses to nutrient limitation can vary significantly within a species and that changes in trait expression may be modified by other environmental variables. These results should be incorporated into existing stoichiometric models and used to investigate the eco-evolutionary consequences of consumer phenotypic variation in response to nutritional stress. Author Keywords: ecological stoichiometry, evolution, life-history, nutrient limitation, nutrient metabolism, zooplankton
Phylogeography and Genetic Structuring of Moose (Alces alces) Populations in Ontario, Canada
Moose are an iconic species, known for their large size and impressive antlers. Eight subspecies are classified in circumpolar regions of the planet - four in North America. Two subspecies are similar in shape and size, the north-western moose (Alces alces andersoni) and the eastern moose (Alces alces americana). It was previously believed that these two subspecies meet in northern Ontario. Earlier genetic population studies used a small number of samples from Ontario, primarily in broad studies covering all of North America. A comprehensive genetic study of moose populations in Ontario has not previously been conducted. We examined the genetic diversity and population structure at 10 polymorphic loci using 776 samples from Ontario, as well as outgroups from representative populations – Manitoba/Cape Breton, representing A. a. andersoni, and New Brunswick/Nova Scotia, representing A. a. americana. Results indicated three genetic populations in the province, in north-western Ontario, north-eastern Ontario and south-central Ontario. RST values, compared against both FST and Jost’s D values for phylogenetic analyses, indicated no phylogenetic pattern which suggests no subspeciation present in the province. Population movement patterns in Ontario were studied. Gene flow was estimated using genetic and spatial data. Isolation by distance was only seen within the first distance class of 100 kilometres and then not seen again at further distances, indicating that moose display philopatry. There were very few migrants travelling across the province, with a greater number moving gradually north and west, towards better habitat and food sources. A forensic database in the form of an allele frequency table was created. Three loci showed very low levels of heterozygosity across all three populations. Probability of identity was calculated for the three populations and quantified. Samples with known geographic origins were run against the database to test for sensitivity, with identification of origin occurring at an accuracy level between 87 and 100%. Within Ontario, there are not two different subspecies, as previously believed, but two different populations of the same subspecies meeting in northern Ontario. The genetic data does not support previous research performed in Ontario. The sample sizes in our research also provide a more comprehensive view of the entire province not seen in any previous studies. The comprehensive research enabled the building of a reliable forensic database that can be used for both management and forensic purposes for the entire province. Author Keywords: Alces alces, Genetic Diversity, Moose, Ontario, Phylogeography, Subspecies
Evaluating the Effects of Habitat Loss and Fragmentation on Canada Lynx
Current major issues in conservation biology include habitat loss, fragmentation and population over-exploitation. Animals can respond to landscape change through behavioural flexibility, allowing individuals to persist in disturbed landscapes. Individual behaviour has only recently been explicitly included in population models. Carnivores may be sensitive to changing landscapes due to their wide-ranging behaviour, low densities and reproductive rates. Canada lynx (Lynx canadensis) is a primary predator of snowshoe hares (Lepus americanus). Both species range throughout the boreal forests of North America, however lynx are declining in the southern range periphery. In this dissertation, I developed new insights into the effects of habitat loss and fragmentation on lynx. In Chapter 2, I created a habitat suitability model for lynx in Ontario and examined occurrence patterns across 2 regions to determine if habitat selection is flexible when different amounts of habitat are available. Although lynx avoided areas with <30% suitable habitat where suitable land cover is abundant, I found that they have flexible habitat selection patterns where suitable land cover is rare and occurred in low habitat areas. In Chapter 3, I investigated the effects of dispersal plasticity on occupancy patterns using a spatially explicit individual-based model. I showed that flexible dispersers, capable of crossing inhospitable matrix, had higher densities and a lower risk of patch extinction. In contrast, inflexible dispersers (unable to cross inhospitable matrix), were most limited by landscape connectivity, resulting in a high extinction risk in isolated patches. I developed three predictions to be explored with empirical data; (1) dispersal plasticity affects estimates of functional connectivity; (2) variation in dispersal behaviour increases the resilience of patchy populations; and (3) dispersal behaviour promotes non-random distribution of phenotypes. Finally, in Chapter 4, I examined the consequences of anthropogenic harvest on naturally cycling populations. I found that harvest mortality can exacerbate the effects of habitat fragmentation, especially when lynx densities are low. Dynamic harvest regimes maintained lynx densities and cycle dynamics while reducing the risk of population extinction. These results suggest that lynx display some flexibility to changing landscapes and that the metapopulation structure is more resilient to increasing habitat loss and fragmentation than previously understood. Future studies should focus on determining a threshold of connectivity necessary for population persistence and examining the effects of habitat loss on the fecundity of lynx. Author Keywords: Fluctuating Populations, Habitat Fragmentation, Landscape Ecology, Occupancy Dynamics, Population Ecology, Spatially Explicit Population Models
Constraints on phenotypic plasticity in response to predation risk
Inducible defenses are plastic responses by an organism to the perception of predation risk. This dissertation focuses on three experiments designed to test the hypothesis that plastic ability is limited by energetic constraints. Chapter 1 provides a general introduction to phenotypic plasticity research and the theoretical costs and limitations affecting the expression of plastic traits. In Chapter 2, I tested the hypothesis that costs of early plasticity may be manifested by a reduced response to risk in later life stages. I found that amphibian embryos are able to detect and respond to larval predators, but that the energetic cost of those plastic responses are not equivalent among behavioural, growth, and morphological traits, and their expression differs between closely-related species. Chapter 3 explicitly examines the relationship between food resource availability and plasticity in response to perceived predation risk during larval development. Food-restricted tadpoles showed limited responses to predation risk; larvae at food saturation altered behaviour, development, and growth in response to predation risk. Responses to risk varied through time, suggesting ontogeny may affect the deployment of particular defensive traits. Chapter 4 examines the influence of maternal investment into propagule size on the magnitude of the plastic responses to predation risk in resulting offspring. I found that females in better body condition laid larger eggs and that these eggs, in turn, hatched into larvae that showed greater morphological plasticity in response to predation risk. Maternal investment can therefore affect the ability of offspring to mount morphological defenses to predation risk. Last, Chapter 5 provides a synthesis of my research findings, identifying specific factors constraining the plastic responses of prey to perceived predation risk. Overall, I found constraints on plastic responses imposed by the current environment experienced by the organism (resource availability), the prior experience of the organism (predator cues in the embryonic environment), and even the condition of the previous generation (maternal body condition and reproductive investment). Together, these findings both provide new knowledge and create novel research questions regarding constraints limiting phenotypic variation in natural populations. Author Keywords: behaviour, inducible defense, Lithobates pipiens, morphometrics, phenotypic plasticity, predation risk
Demography and habitat selection of Newfoundland caribou
The objective of this thesis is to better understand the demography and habitat selection of Newfoundland caribou. Chapter 1 provides a general introduction of elements of population ecology and behavioural ecology discussed in the thesis. In Chapter 2, I examine the causes of long-term fluctuations among caribou herds. My findings indicate that winter severity and density-dependent degradation of summer range quality offer partial explanations for the observed patterns of population change. In Chapter 3, I investigate the influence of climate, predation and density-dependence on cause-specific neonate survival. I found that when caribou populations are in a period of increase, predation from coyotes and bears is most strongly influenced by the abiotic conditions that precede calving. However, when populations begin to decline, weather conditions during calving also influenced survival. I build on this analysis in Chapter 4 by determining the influence of climate change on the interplay between predation risk and neonate survival. I found that the relative equilibrium between bears and coyotes may not persist in the future as risk from coyotes could increase due to climate change. In Chapter 5, I investigate the relationships in niche overlap between caribou and their predators and how this may influence differential predation risk by affecting encounter rates. For coyotes, seasonal changes in niche overlap mirrored variation in caribou calf risk, but had less association with the rate of encounter with calves. In contrast, changes in niche overlap during the calving season for black bears had little association with these parameters. In Chapter 6, I examine broad-level habitat selection of caribou to study trade-offs between predator avoidance and foraging during the calving season. The results suggest that caribou movements are oriented towards increased access to foraging and the reduction of encounter risk with bears, and to a lesser extent, coyotes. Finally, I synthesize the major findings from this thesis and their relevance to caribou conservation in Chapter 7, to infer that Newfoundland caribou decline is ultimately driven by extrinsic and intrinsic elements related to density-dependence. Reduction in neonate survival emerged from nutritionally-stressed caribou females producing calves with lower survival. Author Keywords: Behavioural ecology, Black bear (Ursus americanus), Coyote (Canis latrans), Population ecology, Predator-prey interactions, Woodland caribou (Rangifer tarandus)

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